Late Cretaceous – Holocene, 66–0 Ma
|Crested crane, Balearica regulorum|
Some 5-10 living, see article text.
|Global distribution of the cranes and allies.|
Traditionally, a number of wading and terrestrial bird families that did not seem to belong to any other order were classified together as Gruiformes. These include 14 species of large cranes, about 145 species of smaller crakes and rails, as well as a variety of families comprising one to three species, such as the Heliornithidae, the limpkin, or the trumpeters. Other birds have been placed in this order more out of necessity to place them somewhere; this has caused the expanded Gruiformes to lack distinctive apomorphies. Recent studies indicate that these "odd Gruiformes" are if at all only loosely related to the cranes, rails, and relatives ("core Gruiformes").
There are only two suprafamilial clades (natural groups) among the birds traditionally classified as Gruiformes. Rails (Rallidae), flufftails (Sarothruridae), finfoots and sungrebe (Heliornithidae), adzebills (Aptornithidae), trumpeters (Psophiidae), limpkin (Aramidae), and cranes (Gruidae) compose the suborder Grues and are termed "core-Gruiformes" (Fain et al. 2007). These are the only true Gruiformes. The suborder Eurypygae includes the kagu (Rhynochetidae) and sunbittern (Eurypygidae). These are not even remotely related to Grues. The families of mesites or roatelos (Mesitornithidae), button-quails (Turnicidae), Australian plains-wanderer (Pedionomidae), seriemas (Cariamidae), and bustards (Otididae) each represent distinct and unrelated lineages. Many families known only from fossils have been assigned to the Gruiformes, e.g., Ergilornithidae, Phorusrhacidae, Messelornithidae, Eogruidae, Idiornithidae, Bathornithidae, to name just a few (see below). Though some of these are superficially 'crane-like' and the possibility that some may even be related to extant families traditionally included in the Gruiformes, there are no completely extinct families that can be confidently assigned to core-Gruiformes.
The traditional order Gruiformes was established by the influential German avian comparative anatomist Max Fürbringer (1888). Over the decades, many ornithologists suggested that members of the order were in fact more closely related to other groups (reviewed by Olson 1985, Sibley and Ahlquist 1990). For example, it was thought that sunbittern might be related to herons and that seriemas might be related to cuckoos. Olson and Steadman (1981) were first to correctly disband any of the traditional Gruiformes. They recognized that the Australian plains-wanderer (family Pedionomidae) was actually a member of the shorebirds (order Charadriiformes) based on skeletal characters. This was confirmed by Sibley and Ahlquist (1990) based on DNA–DNA hybridization and subsequently by Paton et al. (2003), Paton and Baker (2006) and Fain and Houde (2004, 2006). Sibley and Ahlquist furthermore removed button-quails (Turnicidae) from the Gruifomes based on large DNA–DNA hybridization distances to other supposed Gruiformes. However, it was not until the work of Paton et al. (2004) and Fain and Houde (2004, 2006) that the correct placement of buttonquails within the shorebirds (order Charadriiformes) was documented on the basis of phylogenetic analysis of multiple genetic loci. Using 12S ribosomal DNA sequences, Houde et al. (1997) were the first to present molecular genetic evidence of gruiform polyphyly, although apparently they were not convinced by it. However, on the basis of numerous additional sequence data, it has been shown decisively that the traditionally recognized Gruiformes consist of five to seven unrelated clades (Fain and Houde 2004, Ericson et al. 2006, Hackett et al. 2008).
Fain and Houde (2004) proposed that Neoaves are divisible into two clades, Metaves and Coronaves, although it has been suggested from the start that Metaves maybe paraphyletic (Fain and Houde 2004, Ericson et al. 2006, Hackett et al. 2008). Sunbittern, kagu, and mesites all group within Metaves but all the other lineages of "Gruiformes" group either with a collection of waterbirds or landbirds within Coronaves. This division has been upheld by the combined analysis of as many as 30 independent loci (Ericson et al. 2006, Hackett et al. 2008), but is dependent on the inclusion of one or two specific loci in the analyses. One locus, i.e., mitochondrial DNA, contradicts the strict monophyly of Coronaves (Morgan-Richards et al. 2008), but phylogeny reconstruction based on mitochondrial DNA is complicated by the fact that few families have been studied, the sequences are heavily saturated (with back mutations) at deep levels of divergence, and they are plagued by strong base composition bias.
The kagu and sunbittern are one another's closest relatives. It had been proposed (Cracraft 2001) that they and the recently extinct adzebills (family Aptornithidae) from New Zealand constitute a distinct Gondwanan lineage. However, sunbittern and kagu are believed to have diverged from one another long after the break-up of Gondwanaland and the adzebills are in fact members of the Grues (Houde et al. 1997, Houde 2009). The seriemas and bustards represent distinct lineages within neoavian waterbirds.
- Family †Parvigruidae Mayr 2005
- Genus †Parvigrus Mayr 2005
- Genus †Rupelrallus Fischer 1997
- Family †Songziidae Hou 1990
- Genus †Songzia Hou 1990
- Suborder Grui
- Super Family †Ergilornithioidea
- Family †Eogruidae Wetmore 1934
- Genus †Sonogrus Kuročkin 1981
- Genus †Eogrus Wetmore 1932 [Progrus Bendukidze 1971]
- Family †Ergilornithidae – may belong in Eogruidae
- Genus †Ergilornis Kozlova 1960
- Genus †Amphipelargus Lydekker 1891 [Urmiornis Harrison 1981]
- Family †Eogruidae Wetmore 1934
- Super Family Gruoidea Vigors 1825
- Family Aramidae Bonaparte 1854 (limpkin)
- Genus †Badistornis Wetmore 1940
- Genus Aramus Vieillot 1816 [Courlili Buffon 1781; Notherodius Wagler 1827] (limpkins)
- Family Psophiidae Bonaparte 1831 (trumpeters)
- Genus Psophia Linnaeus 1758
- Family Gruidae (cranes)
- Genus †Camusia Seguí 2002
- Subfamily Balearicinae Brasil 1913 (crowned cranes
- Genus †Aramornis Wetmore 1926
- Genus †Geranopsis Lydekker 1871
- Genus †Eobalearica Gureev 1949
- Genus Balearica Brisson 1760 [Geranarchus Gloger 1842] (crowned cranes)
- Subfamily Gruinae Vigors 1825
- Genus †"Grus" conferta Miller & Sibley 1942 [Olson & Rasmussen 2001]
- Genus †"Probalearica" mongolica Kurochkin 1985
- Genus †Palaeogrus Portis 1885 [Palaeogrus Salvadori 1884 nomen nudum]
- Genus Grus Brisson 1760 non Moehring 1758 rej. work [Anthropoides Vieillot 1816; Bugeranus Gloger 1841; Megalornis Gray 1841; Leucogeranus Bonaparte 1855; Antigone Reichenbach 1853; Mathewsena Iredale 1914; Mathewsia Iredale 1911; Limnogeranus Sharpe 1893; Laomedontia Reichenbach 1852; Philorchemon Gloger 1842; Scops Gray 1840 non Moehring 1758 rej. work non Bruennich 1772 Savigny 1809] (cranes)
- Family Aramidae Bonaparte 1854 (limpkin)
- Super Family †Ergilornithioidea
- Suborder Ralli
- Family †Aptornithidae (adzebills) (prehistoric) – nested within Rallidae
- Family Sarothruridae (flufftails)
- Genus Rallicula Schlegel 1871 [Corethruropsis Salvadori 1876] (forest-rails)
- Genus Canirallus Bonaparte 1856 [Mentocrex Peters 1933] (pale-throated/wood rails)
- Genus Sarothrura Heine 1890 non Hasselt 1823 [Corethrura Reichenbach 1849 non Hope 1843 non Gray 1846; Daseioura Penhallurick 2003] (flufftails)
- Family Heliornithidae Gray 1841 (finfoots and sungrebe)
- Family Rallidae (crakes, moorhens, gallinules, and rails)
- Genus †Aletornis Marsh 1872 [Protogrus]
- Genus †Australlus Worthy & Boles 2011
- Genus †Baselrallus De Pietri & Mayr 2014
- Genus †Belgirallus Mayr & Smith 2001
- Genus †Capellirallus Falla 1954 (snipe-billed rail)
- Genus †Creccoides Shufeldt 1892
- Genus †Eocrex Wetmore 1931
- Genus †Euryonotus Mercerat 1897
- Genus †Fulicaletornis Lambrecht 1933
- Genus †Hovacrex Brodkorb 1965 (Hova gallinule)
- Genus †Ibidopsis Lydekker 1891
- Genus †Latipons Harrison & Walker 1979
- Genus †Miofulica Lambrecht 1933
- Genus †Miorallus Lambrecht 1933
- Genus †Nesophalaris Brodkorb & Dawson 1962
- Genus †Nesotrochis Wetmore 1918 (West Indian cave-rails)
- Genus †Palaeoaramides Lambrecht 1933
- Genus †Palaeorallus Wetmore 1931
- Genus †Paraortygometra Lambrecht 1933
- Genus †Parvirallus Harrison & Walker 1979
- Genus †Pastushkinia Zelenkov 2013
- Genus †Quercyrallus Lambrecht 1933
- Genus †Rallicrex Lambrecht 1933
- Genus †Rhenanorallus Mayr 2010
- Genus †Vitirallus Worthy 2004 (Viti Levu rails)
- Genus †Wanshuina Hou 1994
- Genus †Youngornis Yeh 1981
- Genus †Rallidae gen. et sp. indet. [Fulica podagrica (partim)] (Barbados rail)
- Genus †Rallidae gen. et sp. indet. (Easter Island rail)
- Genus †Rallidae gen. et sp. indet. (Fernando de Noronha rail)
- Genus †Rallidae gen. et sp. indet. (Tahitian "goose”)
- Genus †Rallidae gen. et sp. indet. (Bokaak "bustard")
- Genus †Rallidae gen. et sp. indet. ('Amsterdam Island' rail)
- Genus Rougetius Bonaparte 1856 (Rouget's Rails)
- Subfamily Rallinae Rafinesque 1815
- Genus †Pleistorallus Worthy 1997 (Fleming's rails)
- Genus Anurolimnas Sharpe 1893 (Chestnut-headed Crakes)
- Genus Biensis (Madagascan Rails)
- Genus Rallus Linnaeus 1758 [†Epirallus Miller 1942]
- Genus †Aphanapteryx von Frauenfeld 1868 [Pezocrex Hachisuka 1953] (Mauritius/Red rails)
- Genus †Erythromachus Milne-Edwards 1873 (Rodriquez rails)
- Genus Dryolimnas Sharpe 1893
- Genus Crex Bechstein 1803 [Crecopsis Sharpe 1893] (greater crakes)
- Genus Lewinia Gray 1855 [Aramidopsis Sharpe 1893; Donacias Heine & Reichenow 1890; Hyporallus Iredale & Mathews 1926]
- Genus Gymnocrex Salvadori 1875 (bare-faced rails)
- Genus Gallirallus Lafresnaye 1841 [Tricholimnas Sharpe 1893; Nesoclopeus Peters 1932; Cabalus Hutton 1874; Habropteryx Stresemann 1932; Eulabeornis Gould 1844; †Diaphorapteryx Forbes 1893; Hypotaenidia Reichenbach 1853; Sylvestrornis Mathews 1928]
- Subfamily Gallinulinae Gray, 1840
- Tribe Pardirallini LIVEZEY, 1998 [Aramidinae] (Wood-rails & allies)
- Tribe Gallinulini GRAY, 1840 [Fulicarinae (Nitzsch 1820) sensu Livezey 1998]
- Genus Pareudiastes Hartlaub & Finsch 1871 [Edithornis] (Woodhens)
- Genus Tribonyx Du Bus de Gisignies 1840 [Brachyptrallus Lafresnaye 1840; Microtribonyx Sharpe 1893] (native-hens)
- Genus Porzana Vieillot 1816 [Limnobaenus Sundevall 1872; Phalaridion Kaup 1829; Porzanoidea Mathews 1912; Porzanoides Condon 1975; Rallites Pucheran 1845; Schoenocrex Roberts 1922; Porphyriops Pucheran 1845]
- Genus Paragallinula Sangster, García-R & Trewick 2015 (Lesser Moorhen)
- Genus Gallinula Brisson 1760 [Hydrogallina Lacépède 1799; Stagnicola Brehm 1831; (Porphyriornis) Allen 1892]
- Genus Fulica Linnaeus 1758 [†Palaeolimnas Forbes 1893]
- Subfamily Porphyrioninae Reichenbach 1849
- Tribe Porphyrionini Reichenbach 1849 (Purple gallinules & swamphens)
- Tribe Himantornithini Bonaparte 1856 (Bush-hens & Waterhens)
- Genus Himantornis Hartlaub 1855 (Nkulenga rails)
- Genus Megacrex D'Albertis & Salvadori 1879 (New Guinea Flightless Rails)
- Genus Aenigmatolimnas (Striped Crakes)
- Genus Poliolimnas Sharpe 1893 (White-browed Crake)
- Genus Gallicrex Blyth 1852 [Gallinulopha Bonaparte 1854; Hypnodes Reichenbach 1853] (Watercocks)
- Genus Amaurornis Reichenbach 1853 [Erythra Reichenbach 1853; Pisynolimnas Heine & Reichenow 1890; Poliolimnas Sharpe 1893] (Bush-hen)
- Tribe Zaporniini Des Murs, 1860 (Old world crakes)
- Tribe Laterallinae TIF 2014 (New world crakes)
- Genus Micropygia Bonaparte 1856 (Ocellated Crakes)
- Genus Rufirallus (russet-crowned crake)
- Genus Laterallus Gray 1855 (ruddy crakes)
- Genus Coturnicops Gray 1855 (barred-backed crakes)
- Genus Hapalocrex (Yellow-breasted Crakes)
- Genus Limnocrex
- Genus Mundia Bourne, Ashmole & Simmons 2003 (Ascension Island Crakes)
- Genus Creciscus Cabanis 1857 [Atlantisia Lowe 1923] (blackish crakes)
When considered to be monophyletic, it was assumed that Gruiformes was among the more ancient of avian lineages. The divergence of "gruiforms" among "Metaves" and "Coronaves" is proposed to be the first divergence among Neoaves, far predating the Cretaceous–Paleogene extinction event c. 66 mya(Houde 2009). No unequivocal basal gruiforms are known from the fossil record. However, there are several genera that are not unequivocally assignable to the known families and that may occupy a more basal position:
- Propelargus (Late Eocene/Early Oligocene of Quercy, France) – cariamid or idornithid
- Rupelrallus (Early Oligocene of Germany) – rallid? parvigruid?
- Badistornis (Brule Middle Oligocene of Shannon County, Missouri) – aramid?
- Probalearica (Late Oligocene? – Middle Pliocene of Florida, France?, Moldavia and Mongolia) – gruid? A nomen dubium?
- "Gruiformes" gen. et sp. indet. MNZ S42623 (Bathans Early/Middle Miocene of Otago, New Zealand) – Aptornithidae?
- Aramornis (Sheep Creek Middle Miocene of Snake Creek Quarries, U.S.) – gruid? aramid?
- Euryonotus (Pleistocene of Argentina) – rallid?
- Family †Gastornithidae (diatrymas) (fossil)
- Family †Messelornithidae (Messel-birds)
- Family †Salmilidae (fossil) – distinct order (Cariamiformes)
- Family †Geranoididae (fossil) – distinct order (Cariamiformes); however, Mayr (2016) argued they might be members of Gruiformes, specifically stem group representatives of the Gruoidea.
- Family †Bathornithidae (fossil) – distinct order (Cariamiformes)
- Family †Idiornithidae (fossil) – distinct order (Cariamiformes)
- Family †Phorusrhacidae (terror birds) (fossil) – distinct order (Cariamiformes)
- Family Cariamidae (seriemas) – Neoavian landbirds – distinct order (Cariamiformes)
- Family Otididae (bustards) – Neoavian waterbirds – distinct order
- Family Eurypygidae (sunbittern) – prospective "Metaves" – new order Eurypygiformes together with kagu
- Family Rhynochetidae (kagu) – prospective "Metaves" – new order Eurypygiformes together with sunbittern
- Family Mesitornithidae (mesites, roatelos, monias) prospective "Metaves" – distinct order
- Horezmavis (Bissekty Late Cretaceous of Kyzyl Kum, Uzbekistan)
- Telmatornis (Navesink Late Cretaceous?)
- Amitabha (Bridger middle Eocene of Forbidden City, Wyoming) – rallid?
- Eobalearica (Ferghana Late? Eocene of Ferghana, Uzbekistan) – gruid?
- "Phasianus" alfhildae (Washakie B Late Eocene of Haystack Butte, U.S.)
- Talantatos (Late Eocene of Paris Bain, France)
- Telecrex (Irdin Manha Late Eocene of Chimney Butte, China) – rallid?
- Neornithes incerta sedis (Late Paleocene/Early Eocene of Ouled Abdoun Basin, Morocco)
- Aminornis (Deseado Early Oligocene of Rio Deseado, Argentina) – aramid?
- Loncornis (Deseado Early Oligocene of Rio Deseado, Argentina) – aramid?
- Riacama (Deseado Early Oligocene of Argentina)
- Smiliornis (Deseado Early Oligocene of Argentina)
- Pseudolarus (Deseado Early Oligocene – Miocene of Argentina) – gruiform?
- Gnotornis (Brule Late Oligocene of Shannon County, Missouri) – aramid?
- Anisolornis (Santa Cruz Middle Miocene of Karaihen, Argentina) – aramid?
- Occitaniavis – cariamid or idiornithid, includes Geranopsis elatus
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